identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
692CB528FFB4FFE7FCD3FF696634FE25.text	692CB528FFB4FFE7FCD3FF696634FE25.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendrobaena Eisen 1874	<div><p>The genus Dendrobaena</p><p>The majority of our specimens belonged to the genus Dendrobaena, which includes many endemic species found in the Caucasus (Kvavadze, 1985; Pizl, 1984; Vsevolodova-Perel, 2003). The monophyly of Dendrobaena was contested: Csuzdi (1984) made a thorough revision of the genus and proposed four species groups: “ octaedra,” “ schmidti,” “ byblica,” and “ veneta .” The conclusions of Csuzdi (1984) were based on the morphology of the male genital pores, the circulatory system, calciferous glands, and muscles. Molecular data appeared to corroborate the division of the genus into several groups. For example, on the phylogenetic reconstructions of Marchán et al. (2021a), representatives of the genus Dendrobaena were scattered throughout the trees, so Marchán et al. (2022) concluded that Dendrobaena formed “several unrelated genus-level clades.”</p><p>Other molecular studies yielded similar results. The majority of them identified a clade consisting of Dendrobaena, Healyella, Spermophorodrilus, and Fitzingeria (Domínguez et al., 2015; Pérez-Losada et al., 2015; Csuzdi et al., 2017; Marchán et al., 2022). In these studies, D. byblica was also found to be outside the main Dendrobaena clade, as in the trees of Marchán et al. (2021a) and on our data (Fig. 1). Three of the aforementioned studies (Domínguez et al., 2015; Pérez-Losada et al., 2015; Marchán et al., 2022) identified Dendrobaena as the sister group to Octolasion + Octodrilus . However, in our study, we found that O. tyrtaeum was more closely related to A. jassyensis rather than to Dendrobaena . We should note that the current analysis includes only one species of Octolasion and no representatives of Octodrilus .</p><p>Previous molecular studies included only a few representatives of Dendrobaena . Here, we added more specimens, and it turns out that all of the specimens except for D. byblica formed a clade. Previous papers also indicated that this taxon was not closely related to other Dendrobaena species (Csuzdi et al., 2017; Domínguez et al., 2015; Latif et al., 2020; Marchán et al., 2022). However, it is important to note that D. byblica is a polymorphic species with multiple cryptic species and a complex taxonomic history (Szederjesi et al., 2018a, 2018b).</p><p>Marchán et al. (2022) suggested that D. byblica might be better classified within the genus Omodeoia Kvavadze (1993) . Previously, Kvavadze (1993) discovered that certain species of the genus Dendrobaena possessed three-striatal genital setae, in contrast to the typical four-striatal structure found in the rest of the genus (Kvavadze, 1993; Kvavadze et al., 2007). Based on this observation, he proposed the genus Omodeoia, with D. byblica designated as the type species. However, limited DNA data is currently available for other potential members of this genus.</p><p>Two early-branching species within Dendrobaena were D. octaedra and D. attemsi . Morphologically, D. attemsi is close to the Balkan species ( D. mahnerti, D. epirotica, D. pindonensis, and D. rhodopensis) (Rota, Erséus, 1997), but based on the position of the clitellum and tuberculae pubertatis, it is close to D. octaedra . Both species are epigeic. D. octaedra is a peregrine species. D. attemsi has a more limited range. D. octaedra is widespread, up to high latitudes in Eurasia and North America (Cameron et al., 2008; Shekhovtsov et al., 2018; Tiunov et al., 2006), while D. attemsi is more common in southern Europe.</p></div>	https://treatment.plazi.org/id/692CB528FFB4FFE7FCD3FF696634FE25	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shekhovtsov, S. V;Rapoport, I. B.;Leonov, P. A.;Poluboyarova, T. V.;Yurlova, G. V.;Shipova, A. A.;Vasiliev, G. V.;Golovanova, E. V.;Roohi Aminjan, A.;Latif, R.	Shekhovtsov, S. V, Rapoport, I. B., Leonov, P. A., Poluboyarova, T. V., Yurlova, G. V., Shipova, A. A., Vasiliev, G. V., Golovanova, E. V., Roohi Aminjan, A., Latif, R. (2024): Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms Diversity & Evolution 24 (4): 477-487, DOI: 10.1007/s13127-024-00656-9, URL: https://doi.org/10.1007/s13127-024-00656-9
692CB528FFB5FFE7FCD3FE376118F9B7.text	692CB528FFB5FFE7FCD3FE376118F9B7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendrobaena hortensis (Michaelsen 1890)	<div><p>D. hortensis group</p><p>A separate clade on the tree (Fig. 1) was formed by D. veneta, D. hortensis, and a putative new species from the Western Caucasus. The affinity between D. veneta and D. hortensis was supported in other studies (Csuzdi &amp; Zicsi, 2003; Kvavadze, 1985). Briones et al. (2009) remarked that D. veneta and D. hortensis are similar in the position of the clitellum and tuberculae pubertatis, chromosome number, and by the absence of diverticules of the calciferous gland in the 10th segment. However, at the same time, they differ in the distance between the setae, the number of spermathecae, and the morphology of the typhlosole, body size, and ecological preferences.</p><p>The prospective new species found in the eastern Caucasus could not be assigned to any existing species within Dendrobaena . Its clitellum starts on the 27th segment, as in D. alpina, D. veneta, and D. hortensis, and ends on 32nd, as in D. schmidti . The position and shape of the tuberculae pubertatis are also similar to D. schmidti . The new species is frequently found under the bark of fallen trees in the arid Eastern Caucasus, just like D. veneta . Dendrobaena sp. probably belongs to the “ alpina ” rather than the “ veneta ” group due to the presence of large calciferous diverticulae in the 11th segment (C. Csuzdi, personal communication). The affinity of Dendrobaena sp. to D. veneta and D. hortensis confirms the complexities that arise during phylogenetic reconstructions.</p><p>This group clustered together with a specimen identified as Dendrodriloides grandis . The genus Dendrodriloides was isolated by Kvavadze (2000) from the genus Eisenia, primarily in the form of the genital setae, as well as the position of the locomotor setae and spermathecae. Species of Dendrodriloides and the closely related genus Dendrobaena typically exhibit similar muscle types and nephridial morphology (Kvavadze, 2000; Zhgenti et al., 2006).</p></div>	https://treatment.plazi.org/id/692CB528FFB5FFE7FCD3FE376118F9B7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shekhovtsov, S. V;Rapoport, I. B.;Leonov, P. A.;Poluboyarova, T. V.;Yurlova, G. V.;Shipova, A. A.;Vasiliev, G. V.;Golovanova, E. V.;Roohi Aminjan, A.;Latif, R.	Shekhovtsov, S. V, Rapoport, I. B., Leonov, P. A., Poluboyarova, T. V., Yurlova, G. V., Shipova, A. A., Vasiliev, G. V., Golovanova, E. V., Roohi Aminjan, A., Latif, R. (2024): Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms Diversity & Evolution 24 (4): 477-487, DOI: 10.1007/s13127-024-00656-9, URL: https://doi.org/10.1007/s13127-024-00656-9
692CB528FFB5FFE7FF6BFDDB605EFE40.text	692CB528FFB5FFE7FF6BFDDB605EFE40.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dendrobaena schmidti (Michaelsen 1907)	<div><p>Dendrobaena schmidti</p><p>A separate clade within the genus Dendrobaena was found to contain D. schmidti and associated taxa (Fig. 1). D. schmidti was initially described as a highly variable species by Michaelsen (1910), who distinguished several morphological forms with subtle differences between them. Pizl (1984) and Kvavadze (1985, 1999, 2000; Kvavadze et al., 1989, 2007) isolated several novel taxa within this complex. Currently, several taxa are known to possess the diagnostic position of the clitellum and tuberculae pubertatis characteristic for D. schmidti: D. baksanensis Pizl (= D. schmidti colchica Kvavadze, 1985), D. jaloniensis (= D. schmidti jaloniensis Kvavadze, 1985), D. surbiensis Kvavadze, 1985 (= D. schmidti surbiensis Kvavadze, 1985), D. malevichi Kvavadze, 1985 (= D. schmidti malevichi Kvavadze, 1985), D. marinae Kvavadze, 1985 (= D. schmidti marinae Kvavadze, 1985), D. adaiensis (Michaelsen, 1900) (= D. schmidti montana (Michaelsen, 1910)), as well as two species initially described as separate species: D. mamissonica Kvavadze, 1985 and Dendrobaena verihemiandra Kvavadze, Patsiashvili &amp; Suladze, 1989 . These taxa differ in body color, the number of spermathecae, the presence or absence of diverticules of calciferous glands, and the number of grooves on genital setae. The validity of these taxa was a matter of debate (Csuzdi et al., 2006; Omodeo &amp; Rota, 1989; Vsevolodova-Perel, 2003, etc.). Vsevolodova-Perel (2003) did not accept the changes proposed by Kvavadze (1985), but recognized D. tellermanica as a separate species with a longer clitellum and a different shape of tuberculae pubertatis. However, our tree does not support this suggestion, i.e., the division of the whole complex into only two species, D. schmidti and D. tellermanica (Fig. 1); D. schmidti appears to be a species complex with high genetic diversity which is often cryptic.</p><p>In addition to the various taxa earlier recognized as related to D. schmidti, the complex also included D. n. nassonovi. This taxon is also a Crimean-Caucasian subendemic, which is believed to have formed in the Greater Caucasus (Perel, 1979). D. n. nassonovi differs from D. schmidti in clitellum position (beginning on the 25th vs. the 26th segment). Also, tuberculae pubertatis of D. n. nassonovi are straight, located on segments 30–32, while those of D. schmidti complex look as two bumps on the 30th and 31st segments. Among the D. schmidti complex, D. baksanensis and D. jaloniensis are the closest to D. n. nassonovi for the number of seminal vesicles and the position of the spermathecae, and D. tellermanica, for the position of the clitellum and the form of tuberculae pubertatis.</p></div>	https://treatment.plazi.org/id/692CB528FFB5FFE7FF6BFDDB605EFE40	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shekhovtsov, S. V;Rapoport, I. B.;Leonov, P. A.;Poluboyarova, T. V.;Yurlova, G. V.;Shipova, A. A.;Vasiliev, G. V.;Golovanova, E. V.;Roohi Aminjan, A.;Latif, R.	Shekhovtsov, S. V, Rapoport, I. B., Leonov, P. A., Poluboyarova, T. V., Yurlova, G. V., Shipova, A. A., Vasiliev, G. V., Golovanova, E. V., Roohi Aminjan, A., Latif, R. (2024): Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms Diversity & Evolution 24 (4): 477-487, DOI: 10.1007/s13127-024-00656-9, URL: https://doi.org/10.1007/s13127-024-00656-9
692CB528FFB5FFE4FCD3F966667AFC6B.text	692CB528FFB5FFE4FCD3F966667AFC6B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Lumbricidae Rafinesque 1815	<div><p>Other Lumbricidae genera</p><p>The rest of Lumbricidae taken into our analysis formed a sister clade to D. byblica . Within this group, one large and well-supported clade included the genera Eisenia, Bimastos, and Rhiphaeodrilus (see below). This clade was weakly related to the group of the Aporrectodea caliginosa complex ( A. caliginosa, A. tuberculata, and A. trapezoides) and Lumbricus ( L. rubellus and L. terrestris). However, on the amino acid trees, the genus Lumbricus moved to the basal position within the D. byblica + rest of the Lumbricidae clade. We hypothesize it could be a long branch attraction artifact; denser sampling within Lumbricus will aid in clarifying its position. The second weakly supported clade contained three groups: A. rosea + A. icterica, P. kaznakovi + H. patriarchalis, and O. tyrtaeum + A. jassyensis . Based on morphological grounds, Octolasion is believed to be related to Aporrectodea based on the similarity of their nephridia, as well as the type of their muscles (Perel, 1979; Vsevolodova-Perel, 1997). Further genomic data from other Octolasion species are required to clarify the affinity between these genera.</p><p>The genus Aporrectodea thus turned out to be polyphyletic, including at least three unrelated groups ( A. caliginosa complex, A. rosea + A. icterica, and A. jassyensis). Other studies yielded similar results for this genus (Domínguez et al., 2015; Pérez-Losada et al., 2015; Csuzdi et al., 2017; Marchán et al., 2022), implying a need for a revision, similar to what was done for Allolobophora (Navarro et al., 2023) . Marchán et al. (2023) suggested that A. rosea, A. icterica, and A. jassyensis are unrelated to the “real” Aporrectodea, but their exact placement is still undefined.</p></div>	https://treatment.plazi.org/id/692CB528FFB5FFE4FCD3F966667AFC6B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shekhovtsov, S. V;Rapoport, I. B.;Leonov, P. A.;Poluboyarova, T. V.;Yurlova, G. V.;Shipova, A. A.;Vasiliev, G. V.;Golovanova, E. V.;Roohi Aminjan, A.;Latif, R.	Shekhovtsov, S. V, Rapoport, I. B., Leonov, P. A., Poluboyarova, T. V., Yurlova, G. V., Shipova, A. A., Vasiliev, G. V., Golovanova, E. V., Roohi Aminjan, A., Latif, R. (2024): Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms Diversity & Evolution 24 (4): 477-487, DOI: 10.1007/s13127-024-00656-9, URL: https://doi.org/10.1007/s13127-024-00656-9
692CB528FFB6FFE5FF6BFC2A666DFC97.text	692CB528FFB6FFE5FF6BFC2A666DFC97.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Eisenia	<div><p>Eisenia + Bimastos + Rhiphaeodrilus clade</p><p>The genus Eisenia was found to be monophyletic (Fig. 1). It was classified into two clades, comprising European and Siberian species, confirming previous research (Shekhovtsov et al., 2020b). This division likely represents an ancient biogeographic divergence. It is worth noting that there is currently insufficient mtDNA data available for Eisenia species from Europe, and no data at all for species from other regions such as Ural or Far Eastern endemics (Perel, 1979).</p><p>Previously, E. lagodechiensis from Georgia was classified under the genus Helodrilus (Michaelsen, 1910) . Vsevolodova-Perel (1997) considered it to be closely related to E. gordejeffi and E. submontana . However, recent studies (Blakemore, 2013) and the Drilobase database now treat it as a subspecies of E. nordenskioldi . However, our results indicate that E. lagodechiensis is a part of the E. fetida / E. andrei complex. These species are morphologically similar and have the same number and position of seminal vesicles and spermathecae (Vsevolodova-Perel, 1997). E. lagodechiensis has a longer clitellum, which ends two segments further compared to E. fetida and E. andrei, and tuberculae pubertatis end one segment further. In contrast to E. fetida and E. andrei, E. lagodechiensis has calciferous glands in the 12th segment and a musculature of the circular type, while E. fetida has a transitional type of longitudinal musculature. In their natural environment, live specimens of E. lagodechiensis closely resemble E. andrei, but are noticeably larger. They have dark purple pigmentation and similar white spots on the lateral surfaces of segments 9–12. Importantly, our findings are not a result of misidentification, as E. lagodechiensis did not match any of the distinct genetic lineages within this group.</p><p>E. nordenskioldi exhibits extensive morphological and genetic diversity, as shown by various studies (Malevich, 1956; Perel, 1997; Vsevolodova-Perel, 1997; Blakemore, 2013; Shekhovtsov et al., 2020b). Consistent with previous research on mitochondrial genomes (Shekhovtsov et al., 2020a), it was divided into two main clades that grouped together with E. tracta and E. altaica (Fig. 1). These findings reinforce the notion that E. nordenskioldi sensu lato is not a single species, but rather a complex consisting of multiple species from the Altai Mountains and the surrounding region. The Altai Mountains are known for harboring numerous endemic Eisenia species (Vsevolodova-Perel, 1997), making it intriguing to investigate how many of these species would also fall within this clade.</p><p>Previous studies demonstrated that Eisenia, Bimastos, and Eisenoides form a clade (Csuzdi et al., 2017; Marchán et al., 2022). In our study, we also identified a clade consisting of Eisenia, Bimastos, and Rhiphaeodrilus diplotetrathecus . Initially described as Allolobophora handlirschi diplotetratheca Perel, 1967, together with P. kaznkovi and several other species, it was later transferred to the subgenus Svetlovia Perel, 1977 . Representatives of this subgenus were believed to differ from the rest of the genus Allolobophora Eisen, 1873 by the absence of calciferous glands in the diverticulae of the 10th segment, the position of the spermathecae not extending beyond the 10/11 dissepiment, the presence of three to four pairs of seminal vesicles, by weak development of glandular fields around the male genital openings, and long, elongated tuberculae pubertatis stretching almost along the entire clitellum. However, the name Svetlovia turned out to be occupied (genus Svetlovia Chekanovskaya, 1975 within Tubificidae); Easton proposed to elevate the taxon to the genus rank with the name Perelia (Easton, 1983) . Subsequently, the species was transferred to Alpodinaridella (Mrsic, 1987), and finally to the monotypic genus Rhiphaeodrilus (Csuzdi &amp; Pavlíček, 2005) based on three pairs of reddish-violet vesicles and pinnate musculature. Our molecular data support the isolation of R. diplotetrathecus from Perelia, as it was not related to P. kaznakovi on our phylogenetic tree (Fig. 1). Unfortunately, the DNA extracted from our samples of another Ural endemic, P. tuberosa, was highly degraded and could not be sequenced.</p><p>The three genera, Eisenia, Bimastos, and Riphaeodrilus, have a morphological synapomorphy: their nephridial bladders form a forward-bent loop. This distinguishes them from their sister group on the tree ( Lumbricus, Aporrectodea, Helodrilus species), whose nephridial bladders are bent backwards or were lost. Therefore, the form of nephridial bladders appears to be an important diagnostic character, as suggested by Gates (1975) and Perel (1976) (but with some exceptions (Marchán et al., 2021b)).</p><p>Representatives of the genus Bimastos (Dendrodrilus) are believed to be of North American origin (Csuzdi et al., 2017; Gates, 1969). Csuzdi et al. (2017) proposed that their ancestors likely dispersed through the Bering Strait into North America, and more recently, a few species, notably B. parvus and B. rubidus, returned to Eurasia as migratory species. We believe that further data is necessary to fully investigate this hypothesis. However, the affinity between the North American and Ural endemic species is intriguing and may reflect dispersal patterns of ancestral taxa.</p><p>Therefore, this clade represents an interesting example of phylogeographic patterns across a large territory, in contrast to the rest of the Lumbricidae genera and clades, which are mostly native to the Europe or Middle East. Riphaeodrilus is an endemic of the Urals, Bimastos is reportedly native to North America (Csuzdi et al., 2017), and Eisenia has clear phylogeographic patterns with an Altai clade (Shekhovtsov et al., 2020b).</p></div>	https://treatment.plazi.org/id/692CB528FFB6FFE5FF6BFC2A666DFC97	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Shekhovtsov, S. V;Rapoport, I. B.;Leonov, P. A.;Poluboyarova, T. V.;Yurlova, G. V.;Shipova, A. A.;Vasiliev, G. V.;Golovanova, E. V.;Roohi Aminjan, A.;Latif, R.	Shekhovtsov, S. V, Rapoport, I. B., Leonov, P. A., Poluboyarova, T. V., Yurlova, G. V., Shipova, A. A., Vasiliev, G. V., Golovanova, E. V., Roohi Aminjan, A., Latif, R. (2024): Phylogeny of Lumbricidae based on mitochondrial genomes with emphasis on the genus Dendrobaena. Organisms Diversity & Evolution 24 (4): 477-487, DOI: 10.1007/s13127-024-00656-9, URL: https://doi.org/10.1007/s13127-024-00656-9
